Aquatic Insects
Vol. 32, No. 4, December 2010, 265–280
New species and new stage descriptions of Campsurus major species
group (Polymitarcyidae: Campsurinae), with first report of silk-case
construction in mayfly nymphs
Carlos Molineria* and Daniel Emmerichb
a
CONICET, Instituto de Biodiversidad Neotropical, Facultad de Ciencias Naturales e IML, San
Miguel de Tucumán, Argentina; bSección Entomologı´a PEDECIBA, Facultad de Ciencias,
Universidad de la República, Montevideo, Uruguay
(Received 9 September 2009; final version received 31 July 2010)
Campsurus major, C. argentinus and C. amapaensis sp. nov. form a monophyletic
group supported by the following apomorphies in the male genitalia: robust
pedestals with a widely rounded inner-posterior margin and a more acute and
thinner outer margin, penes strongly sclerotised and curved ventrally, each lobe
twisted outwards, and with a small membranous lobe on ventral margin. The
nymphs of Campsurus major and C. argentinus and the female adult of C. major are
described for the first time and illustrated here; the male adults of both species are
redescribed. A new species, C. amapaensis, is described from male imagos. The
following new synonymies are proposed: Campsurus argentinus Esben-Petersen
(¼ C. pallidus Needham & Murphy, junior synonym) and C. major Needham &
Murphy (¼ C. brasiliensis Traver, junior synonym). The nymphs of both species are
reported to use silk to build soft cases and to glue substrate particles. A list of
characters useful to distinguish the treated species from others in the genus is given.
Keywords: Campsurus argentinus; C. brasiliensis; C. pallidus; C. amapaensis;
Ephemeroidea; case-building; silk
Introduction
Polymitarcyidae belongs to a group known as ‘‘burrowing mayflies’’ (Ephemeroidea), characterised by its relatively large nymphs adapted to live in tunnels they
excavate in different substrates (soft mud, hard clay, plant tissues, McCafferty 1975).
The family is known from three major groups: Polymitarcyinae, Asthenopodinae
and Campsurinae (McCafferty 1975; Kluge 2004). Campsurinae, with two genera
and about 50 species, shows a Panamerican distribution, with its main diversity in
South American tropics (Domı́nguez, Molineri, Pescador, Hubbard and Nieto
2006). This paper deals with three species: Campsurus argentinus Esben-Petersen,
1912; C. major Needham & Murphy, 1924; and C. amapaensis sp. nov. We are
describing the nymphs of C. argentinus and C. major and interestingly both were
found to produce and use silk for their protection. In the case of C. argentinus we
could confirm the use of the silk for case construction.
*Corresponding author. Email: [email protected]
ISSN 0165-0424 print/ISSN 1744-4152 online
Ó 2010 Taylor & Francis
DOI: 10.1080/01650424.2010.533131
http://www.informaworld.com
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C. Molineri and D. Emmerich
The aim of the present paper is to describe the previously unknown nymphal
stages of C. argentinus and C. major and their particular way of living. Also, adults
of both species are redescribed and illustrated, and the female imago of C. major is
described for the first time. New country records and synonyms are given for both
species. Lastly, a new species from northeast Brazil is described from male imagos,
Campsurus amapaensis sp. nov.
Materials and methods
Morphological terms in descriptions are mainly from Kluge (2004). SEM
photographs were obtained with a JEOL 35CF scanning electron microscope at
25 kV. Larval parts were dehydrated in a graded ethanol series, dried by critical
point-method, mounted with double-sided tape on SEM stubs and sputter coated
with gold. Depositories: CUIC (Cornell University Insect Collection, Ithaca, NY),
FAMU (Florida A&M University, Tallahassee, FL), IML (Instituto Miguel Lillo,
Tucumán, Argentina), MCR (Museo de Ciencias de la República, Montevideo,
Uruguay), and INPA (Instituto Nacional de Pesquisas da Amazônia, Manaus,
Brazil).
Campsurus argentinus Esben-Petersen, 1912 (Figures 1, 4–18, 35, 36)
Campsurus argentinus Esben-Petersen, 1912: 334 (male); Ulmer, 1920: 106; Lestage, 1923: 120;
Needham & Murphy, 1924: 16; Ulmer, 1942: 117 (male); Traver, 1947: 379; Traver, 1950: 593.
Campsurus pallidus Needham & Murphy, 1924: 20 (male); Traver, 1947: 380 New synonym.
Type material. Campsurus pallidus holotype < imago, slide (CUIC N8 623.1) with genitalia,
head, forelegs, and 1 pair of wings from Perú [sic], rı́o Alto Paraná, 18-Jan-1920. In the
original description type locality data is given as Argentina-Paraguay, above Posadas, 18-I1920. Types of C. argentinus could not be located; identification of the material was done by
comparison with original figures and description.
Additional material. Five < and 1 , (IML) from ARGENTINA: Misiones, P.P. Urugua-ı́, A8
Uruzú, S 258 510 2900 – W 548 100 1000 , 7-11/XII/1999, C. Molineri col. One < imago (IML)
from BRASIL: Rio Grande do Sul, Rio Lageado Grande, 29/I/1982, N. Santos & N.F. Netto
cols. Four < imagos (IML) from URUGUAY: Rivera, Mina de Corrales, A8 Corrales, S 318
340 3400 – W 558 270 5800 , 135 m, 19/II/2008, D. Emmerich & C. Molineri cols.; 4 < and 2 ,
imagos (IML) from URUGUAY: Maldonado, R60 km 23, A8 El Rodeo, S 348 400 2000 – W
558 140 1500 , 75 m, 7/XI/2008, D. Emmerich & C. Molineri cols.; 1 < imago (IML slide 380)
from URUGUAY: Treinta y Tres, Qda. de los Cuervos, S 328 550 27.300 – W 548 270 34.400 , 125
m, 8/XI/2004, light trap, D. Emmerich col.; 1 < subimago (IML slide 400) from URUGUAY:
Cerro Largo, A8 Tacuarı́, S 328 310 5100 – W 548 070 4800 , 92 m, light trap, 20/II/2008, D.
Emmerich & C. Molineri cols.; 7 < and 8 , (MCR) from URUGUAY, Tacuarembó,
Tacuarembó Chico, 24-25/I/1980; 10 < and 10 , imagos (MCR) from URUGUAY,
Sepulturas, rio Cuareim, 15/I/1952, C. Carbonell & L. de Zolessi cols.; 37 nymphs from (IML,
MCR) URUGUAY: Artigas, A8 Catalán Chico, S 308 420 28.800 – W 568 190 17.600 , 128 m, 18/
II/2008, D. Emmerich & C. Molineri cols.; 8 nymphs from URUGUAY: Artigas, Finca
Mataojo Grande, A8 Mataojo Grande (IML, used in SEM study) , S 318 170 29.700 – W 568 130
1000 , 274 m, 18/II/2008, D. Emmerich & C. Molineri cols.; 2 nymphs (MCR) from
URUGUAY: Tacuarembó, R5 km 343, A8 La Qda. Grande, S 328 050 3100 – W 568 060
0700 , 135 m, 22/II/2008, D. Emmerich col.; and 3 nymphs (MCR) from URUGUAY: Rivera,
Santa Ernestina, R 29, A8 sin nombre, S 318 320 23.800 – W 558 330 4200 , 146 m, 19/II/2008,
D. Emmerich & C. Molineri cols.
Figures 1–7. (1) Campsurus argentinus, nymph and silk-case; (2–3) Campsurus major, (2) nymph, general view; (3) head and pronotum, dorsal view.
Campsurus argentinus, imago; (4) male head and pronotum; (5) female head and pronotum; (6) male abdominal terga I–VI, dorsal view; (7) male
abdominal segments III–V, lateral view (td ¼ transversal pale dashes).
Aquatic Insects
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Figures 8–12. Campsurus argentinus; (8–9) male genitalia in (8) ventral view; (9) lateral view;
(10) detail of penes, ventral view; (11) detail of pedestal, ventral view; (12) detail of penes,
lateral view.
Male imago
Length (mm): body, 10.8–11.0; forewing, 9.5–10.3; hind wing, 4.2; cerci, 28.0–30.1;
fore leg, 5.0–5.2. General colouration yellowish white with brownish markings
dorsally.
Head dorsally blackish (Figure 4), scape and pedicel brownish, flagellum hyaline.
Thorax. Pronotum translucent shaded brownish medially and laterally, blackish
medial line well marked (Figure 4). Meso- and metanota shaded black on carinae.
Pleura and sterna yellowish white except on prothorax shaded purplish grey;
mesofurcasternal plates contiguous basally but diverging distally. Forelegs shaded
purplish brown except apical 1/3 of each tarsite, pale; vestiges of middle and hind
legs yellowish white.
Wings (Figures 35, 36). Membrane hyaline, except C and Sc areas tinged with
brown; veins brownish, becoming darker towards fore margin.
Abdomen whitish translucent, shaded with brownish on terga (Figures 6 and 7): a
thin medial line on terga IV–X, a paler band around it extending from terga I–VIII,
and with shading darker more laterally; terga I–II with a pair of submedian small
anterior marks, terga III–VII shaded more widely except on pale lateral transversal
band (td in Figures 6, 7); terga VIII–IX widely pigmented and with small dark and
pale marks; tergum X paler.
Aquatic Insects
269
Figures 13–18. Campsurus argentinus, nymph; (13) general frontal view; (14) head, dorsal
view; (15) foreleg, ventral view; (16) middle legs, ventral view (left), dorsal view (right); (17)
hind leg, dorsal view; (18) abdominal sternum IX.
Genitalia (Figures 8 – 12) whitish yellow, forceps shaded grey, sclerotised apex
of penes orangeish. Penes (Figures 10, 12) strongly curved ventrally, each penial
arm twisted outwards and with its prominent dorsal (posterior) margin very
sclerotised; ventral margin of each penial arm membranous and with a small
cylindrical projection. Pedestals short and somewhat flattened, inner margin well
developed and rounded, outer margin much smaller (Figure 11). Caudal filaments
whitish translucent.
Female imago
Length (mm): body, 10.6–12.7; forewing, 12.6–14.7; hind wing, 5.4–6.3; cerci, 4.6;
fore leg, 1.5–1.7. As in male, exceptions follow.
Head more heavily shaded with black (Figure 5). Foreleg paler, shaded slightly
with purplish. Abdominal terga III–VII with a pale transverse band on lateral areas
clearly indicated, spiracles surrounded with blackish; terga VIII–X more widely and
strongly pigmented. Wing membrane whitish translucent shaded very slightly with
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brown, longitudinal veins brownish, crossveins paler. Abdominal sternum VIII with
a single bottle-like socket on median zone of fore margin, sternum with a grey dash
at the base of this socket.
Mature nymph
Length of male (mm): body, 9.0; terminal filament, 6.0; cerci, 6.5. Length of female
(mm): body, 9.2–13.0; terminal filament, 5.8–6.0; cerci, 6.0–7.5. General colouration
whitish yellow with brownish marks.
Head (Figures 1, 13, 14) shaded brownish dorsally except on whitish genae and
yellowish frontal projection. Mouthparts and antennae whitish except sclerotized
areas, spines and thick setae yellowish to orangeish; mandibular tusks with variable
number of denticles anteriorly to subbasal tubercle (5–10 on left tusk, 6–8 on right
tusk).
Thorax. Pronotum shaded brownish almost in the entire dorsum, darker on
medial line (pale longitudinal line surrounded by black parallel lines at each side)
and other small marks (Figure 1); meso- and metanotum shaded mainly on median
band, carinae darker. Wing buds shaded grey along main veins. Thoracic pleura and
sterna without marks. Legs whitish except at joinings, yellowish. Forelegs with
numerous fringed setae (Figures 13, 15); middle legs (Figure 16) and hind legs
(Figure 17) with long and short simple setae.
Abdomen whitish shaded brownish dorsally with a pattern similar to that
described for the male adult: tergum I shaded widely except on median band; terga
II–VII shaded almost completely except on a long paler transverse band at each side;
terga VIII–X widely shaded except on four submedian pale marks (partially fused in
tergum 10) and an additional pair of sublateral pale marks in tergum VIII.
Abdominal sterna without marks, sterna VIII–X with many long setae, longer on a
median row of sternum IX (Figure 18). Gills (Figure 1): vestigial gill I whitish,
bilamellate with a small and thin ventral lobe and a larger rounded dorsal lobe; gills
II–VII with ventral lobe also smaller than dorsal lobe but increasing in length
posteriorly, so on gill VII both lobes are subequal in length; fringes of gills whitish,
lamellae widely brownish (somewhat paler on ventral lobe). Caudal filaments whitish
covered with thin yellowish setae.
Biology
Mayflies are not widely known as silk producers but some Polymitarcyidae nymphs
have long been reported to produce silk from modified Malpighian tubules
(Hartland-Rowe 1958; Sattler 1967). Nymphs of Asthenopodinae (Asthenopus and
Povilla) use the silk to coat U-tunnels they burrow in different submersed plant
tissues (Hartland-Rowe 1958; Sattler 1967). Campsurinae nymphs (the sister group
of Asthenopodinae) burrow their tunnels in soft substrates like mud or sand
(Campsurus) or in hard-clay river banks (Tortopus). Nymphs of an unidentified
Amazonian species of Campsurus were reported to coat their burrows with silk,
thus preventing their collapse (Sattler 1967). Tortopus, living in more stable
substrates, apparently do not coat their tunnels (Scott, Berner and Hirsch 1959;
Molineri 2008).
The nymphs of C. argentinus were collected in streams with a rocky substrate.
Each nymph was found inside a cylindrical silk case (Figure 1) slightly fixed to the
Aquatic Insects
271
upper surface of rocks, mainly in riffle zones but also in pools near the shore. Cases
are made of a thin layer of silk with fine sediment on the outer surface. Both ends of
the tube are completely open, and cases are large enough to hold the entire nymph
inside. As the cases are very soft and poorly fixed to the rocks, their role seems
related more to a camouflage than to a mechanic protection for the nymphs.
It is probable that the silk is produced in modified Malpighian tubules, as
reported for other species of the genus (Hartland-Rowe 1958; Sattler 1967). It has
been reported that the silk produced in insect Malpighian systems are not delivered
through a specialised spinning organ (Craig 1997), and concordantly in the case of
C. argentinus the silk is apparently liberated from the anus. A mediolongitudinal row
of long setae on abdominal sterna IX (Figure 18) may help during case construction,
but this behaviour was not observed.
Discussion
All three species treated here form a monophyletic group (below ‘‘C. major-group’’)
defined by characters from male genitalia. Domı́nguez et al. (2006) included the species
of the C. major-group in a larger assemblage (C. albifilum-group). Both groups of
species share some synapomorphies in male genitalia (penes with an additional
membranous ventral lobe and pedestals relatively flat. The C. major-group is defined
here by the following apomorphies (characters that can be used to separate the three
species of the group from all other species in the genus): (1) pedestals robust, with a
widely rounded inner-posterior margin and a smaller and thinner outer margin
(Figures 11, 24, 34); (2) penes strongly sclerotised and curved ventrally, each
lobe twisted outwards, and with a small membranous lobe on ventral margin (Figures
8–10, 23–26, 30–33); (3) male gonopore large, clearly visible on the apex of penial lobe,
and opening towards the medial line (Figures 10, 23, 33).
The useful characters to separate C. argentinus from the other two species in the
group are, in the adults
(1) medium size (body 10.8–13.0 mm, forewings 9.5–13.0 mm);
(2) blackish pigments on dorsum of head widely and strongly marked in fresh
material (Figures 4 and 5);
(3) pronotum shaded blackish, including the medial line (Figures 4 and 5);
(4) abdominal colour pattern with clearly pale transverse dashes on terga II–VII
(Figures 6 and 7);
(5) pedestals with both posterolateral margins (inner and outer) approximately
of the same length (Figure 11);
(6) penes relatively slender and moderately twisted (Figures 10 and 12).
In mature nymphs, the above characters (2)–(4) are also useful to distinguish the
species, but character (3) may show variations (pronotum with a thin pale
longitudinal line surrounded by black parallel lines at each side); other characters are
(1) mandibular tusks with variable number of denticles anteriorly to sub-basal
tubercle (5–10 on left tusk, 6–8 on right tusk);
(2) wing buds shaded grey along main veins;
(3) gills II–VII with lamellae widely shaded with brownish, somewhat paler on
ventral lamellae (Figure 1).
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Campsurus pallidus Needham & Murphy was described for one male from Alto
Parana (above the locality of Posadas in Argentina), with a body length of 9 mm and
fore wings of 12 mm (Needham and Murphy 1924, p. 20). Campsurus argentinus
Esben-Petersen (1912) was also described from one male from Alto Parana (between
Posadas and Corrientes), with a length of 13 mm for the body and the fore wing. At
the moment both species are not distinguishable since the genitalia, wing venation,
size range and general colouration coincide completely. For these reasons Campsurus
pallidus Needham & Murphy (1924) is proposed here as a junior subjective synonym
of C. argentinus Esben-Petersen (1912).
Distribution
This species was previously known from Argentina-Paraguay border (Alto Paraná),
and is reported here for the first time from Brazil (Rio Grande do Sul) and
Uruguay.
Campsurus major Needham & Murphy, 1924 (Figures 2, 3, 19–26, 37, 38)
Campsurus major Needham & Murphy, 1924: 17 (male); Traver, 1947: 378.
Campsurus brasiliensis Traver, 1944: 36 (male), new synonym.
Type material. Campsurus major holotype < imago, slides (CUIC N8 618.1) with genitalia and
1 pair of wings from ARGENTINA, Buenos Aires, 16-XII-1898. Five additional slides (also
from CUIC, designated as paratypes by Traver 1947), same data as holotype, representing
three < imagos (genitalia, wings, forelegs and head).
Additional material. Twenty seven < and 6 , imagos (IML) from ARGENTINA: Misiones,
P.N. Iguazú, rio Iguazú, Puerto Canoas, S 258 410 3900 – W 548 260 5100 , 26/XI/1998, E.
Domı́nguez et al. cols.; 1 < imago (IML) from ARGENTINA: Misiones, P.P. Urugua-ı́, A8
Uruzú, S 258 510 2900 – W 548 100 1000 , 7–11/XII/1999, C. Molineri col. Eight < imagos
(FAMU) from BRASIL: Parana, rio dos Patos, 3 km E of Prudentópolis, 16/III/1969 W.L. &
J.G. Peters cols. Eleven < imagos (MCR) from URUGUAY: Paysandu, Santa Rita, XII-1959,
Carbonell col.; 1 < imago (MCR) from URUGUAY: Maldonado, R60 km 23, A8 El Rodeo,
S 348 400 2000 – W 558 140 1500 , 75 m, 7/XI/2008, D. Emmerich col.; 4 < (IML slide 379) from
URUGUAY: Treinta y Tres, Qda. de los Cuervos, S 328 550 27.300 – W 548 270 34.400 , 125 m,
light trap, 8/XI/2004, D. Emmerich col.; 6 < adults (IML) from URUGUAY: Cerro Largo,
A8 Tacuarı́, S 328 310 5100 – W 548 070 4800 , 92 m, light trap, 20/II/2008, D. Emmerich & C.
Molineri cols.; 1 < subimago (IML) from URUGUAY: Durazno, Sarandı́ del Yi, Rı́o Yi, S
338 200 4100 – W 558 370 0700 , 132 m, 22/II/2008, D. Emmerich & C. Molineri cols.; 4 nymphs
(IML, MCR) from URUGUAY, Rivera, Tranqueras, A8 Tacuarembó, S 318 100 4000 – W 558
450 4400 , 147 m, 19/II/2008, D. Emmerich & C. Molineri cols.
Male imago
Length (mm): body, 13.2–16.0; forewing, 12.0–14.2; hind wing, 5.2–6.0; cerci, 32.0–
42.0; foreleg, 5.5–7.2. General colouration yellowish white with very few dark
markings.
Head shaded purplish gray dorsally, stronger between ocelli, a transverse black
line (shallow V-mark) extends between lateral ocelli, posteriorly to this line the
shading is much paler (Figure 19). Antennae: pedicel shaded purplish grey, also
scape but paler.
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273
Figures 19–22. Campsurus major, imago; (19) male head and pronotum; (20) female head
and pronotum; (21) male abdomen, dorsal view; (22) male abdomen, lateral view.
Thorax. Prothorax translucent, diffusely shaded with gray dorsally except on
mediolongitudinal line (Figure 19). Meso- and metathorax yellowish white without
markings. Foreleg shaded rather uniformly with purplish grey; middle and hind legs
yellowish white.
Wings (Figures 37 and 38). Wing membrane hyaline; veins C, Sc & R (and
crossveins therein) shaded brown on basal 2/3, remaining veins yellowish to whitish
translucent.
Abdomen. Pattern similar to that described for the males of C. argentinus but the
pale lateral (or transversal dashes) marks are less distinguishable and blurred
(Figures 21 and 22).
Genitalia (Figures 23 – 26) whitish except pedestals yellowish white, and sclerotised
portions of penes orangeish. Pedestals with rounded inner margin, outer margin short
and acute (Figure 24); penes (Figures 23, 25, 26) very similar to C. argentinus but larger,
each lobe very wide and strongly twisted, the dorsal (posterior margin) strongly
developed; a small membranous projection is also present on ventral margin of each
penes. Caudal filaments whitish translucent.
Female imago
Length (mm): body, 13.8–15.7; forewing, 15.5–17.0; hind wing, 8.0; cerci, 5.3; foreleg, 1.8–2.5. Head colour pattern as in Figure 20. Rest as male except purplish grey
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C. Molineri and D. Emmerich
Figures 23–26. Campsurus major; (23) detail of penes, ventral view; (24) male genitalia,
ventral view; (25–26), male genitalia, lateral view.
marks present on mesonotal carinae; wing membrane whitish translucent; abdomen
shaded very slightly with purplish grey. Abdominal sternum VIII as described for the
previous species.
Nymphs (immature) (Figures 2 and 3)
Head colour pattern similar to male imago (with a black transverse line between
ocelli) (Figure 3); frons and genae not shaded, paler than the rest; with a pair of
occipital submedian star-like greyish marks. Mouthparts and antennae whitish
except sclerotised areas, spines and thick setae yellow to orangeish; all the nymphs
studied (n ¼ 4) show six denticles anteriorly to sub-basal tubercle on left mandibular
tusk and seven denticles on the right tusk.
Thorax. Pronotum shaded diffusely with grey on anterior 2/3 of their length,
mediolongitudinal line paler. Meso- and metanotum with small irregular grey marks
medially and costal veins of wingbuds. Legs whitish with yellowish setae and
articulations.
Abdomen yellowish white shaded brownish submedially on terga I–II, terga III–
VII shaded more widely except on thin transversal pale dashes (broken by
dark pigments at middle); terga VIII–X widely shaded except on four submedian
pale marks (partially fused in tergum X) and an additional pair of sublateral pale
marks in tergum VIII. Abdominal sterna without marks, sternum IX with long
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275
medial setae. Gills II–VII widely shaded with brownish, restricted to trachea in
ventral lamellae (Figure 2). Caudal filaments whitish covered with thin yellowish
setae.
Biology
We only collected four immature nymphs attributable to this species, one of the
nymphs was found inside a hollow stick with some parts glued together by thin silk
filaments. No silk case was observed in this species.
Discussion
The characters useful to separate C. major from the other species of the C. majorgroup are, in the adults
(1) larger size (body 13.2–16.0 mm, forewings 12.0–16.0 mm);
(2) blackish pigments on dorsum of head mainly restricted to a transversal
shallow V-mark between ocelli (Figures 19 and 20);
(3) pronotum shaded greyish to blackish, except on paler medial line (Figures 19
and 20);
(4) abdominal colour pattern weak and not forming completely distinguishable
pale transverse dashes on terga II–VII (Figures 21 and 22);
(5) pedestals with outer posterolateral margin somewhat longer than the inner
one (Figure 24);
(6) penes large, wide, strongly twisted (Figures 23 – 26).
In mature nymphs, the above characters (2) to (4) are also useful to distinguish the
species, and also:
(1) mandibular tusks with six denticles anteriorly to sub-basal tubercle on left
tusk and seven on the right tusk;
(2) wing buds shaded grey on costal margin;
(3) gills II–VII with dorsal lamellae widely shaded with brownish, restricted to
trachea in ventral lamellae (Figure 2).
Campsurus major Needham & Murphy (1924) was described for a single male
from Buenos Aires (16-XII-1898, M. Baer col.), with a body length of 13 mm and
a forewing of 20 mm. Traver (1947) corrected the length of the wing to 16 mm,
and designated three additional male imagos as paratypes. Campsurus brasiliensis
Traver (1944) was described from 42 male imagos from Brazil (Rio Grande do
Sul, Lagoa dos Quadros, XI-1941, H. Kleerekoper col.). The author separated
this species from C. major because this last species presents (1) general
colouration yellowish, (2) in the hind wings the cross veins of C area are better
visible, and (3) there is a network of cross veins in the anal margin of hind wings.
All these characters are variable among the material studied and thus are not
reliable to separate both species. The general colouration is always very pale but
varies from whitish to yellowish even between individuals of the same population.
The hind wings present cross veins in costal area normal to weak, and the cross
veins in anal area may be strongly to not anastomosed at all (even between the
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C. Molineri and D. Emmerich
two wings of the same specimen). For these reasons we here propose Campsurus
brasiliensis Traver as a junior subjective synonym of Campsurus major Needham
& Murphy.
Distribution
Originally described from Buenos Aires (Argentina) and Rio Grande do Sul (Brazil),
we here firstly record this species from Parana (Brazil), NE Argentina (Misiones) and
Uruguay.
Campsurus amapaensis sp. nov. (Figures 27–34, 39–40)
Type material. Holotype (INPA) and paratype (IML) < imagos from BRAZIL:
Amapá, Municipio de Amapá, Cachoeira Grande, 21/XII/2008, light trap, I. Santos Andrade
col.
Male imago
Length (mm): body, 8.5–11.0; forewing, 9.0–9.8; hind wing, 4.3–5.0; cerci broken off
and lost; foreleg, 4.0–4.3. General colouration yellowish white, dark shading on
dorsum of abdomen.
Head shaded purplish grey on the entire dorsum (Figure 27). Antennae with
scape whitish shaded with hyaline.
Thorax. Pronotum yellowish translucent shaded diffusely with grey, darker on
carinae and mediolongitudinal line (Figure 27); propleura and sternum yellowish
white shaded grey more strongly, a blackish mark is present posterolaterally to
coxae I. Mesonotum whitish yellow with a light grey medial mark between
posteroscutal protuberances (PSP), carinae and medial line shaded grey;
mesopleura and sterna whitish yellow with some grey shading on carinae, wing
base and furcasternum. Metathorax whitish yellow with few grey stripes. Foreleg
shaded rather uniformly with dark purplish grey, except at bases of tarsomeres and
claws, slightly paler; middle legs shaded grey almost completely, hind legs shaded
grey more slightly.
Wings (Figures 39 and 40). Wing membrane hyaline except C and Sc spaces
slightly tinged with purplish grey; longitudinal and cross veins light purplish grey;
hind wing with paler veins.
Abdomen (Figures 28 and 29) translucent whitish shaded with grey dorsally,
except on medial line whitish, terga shaded more strongly posteriorly; pattern similar
to the other two species, pale lateral marks on terga II–VII ‘‘broken’’ at the middle,
the medial portion of these marks tending to form well defined submedian pale
circles on terga V–VII; terga VIII–IX with two pairs of small submedian pale marks;
tergum X pale yellow almost without shading.
Genitalia (Figures 30 – 34) whitish except pedestals and apex of penes yellowish;
forceps strongly shaded with black. Pedestals narrow, longer than wide, with
rounded inner margin ending in a somewhat acute point, outer margin completely
reduced (Figure 34); penes (Figures 32 and 33) wide at base becoming thinner
apically, twisted moderately, apex of penes acutely projected; small membranous
projection present on ventral margin of each penes. [Caudal filaments broken off
and lost.]
Aquatic Insects
277
Figures 27–29. Campsurus amapaensis sp. nov., male imago; (27) head and pronotum; (28)
abdominal segments II-X, dorsal view; (29) abdominal segments II–VII, lateral view.
Female imago and nymph unknown.
Discussion
The characters useful to separate C. amapaenis sp. nov. from the other two species of
the C. major-group are, in the adults
(1) small size (body 8.5–11.0 mm, forewings 9.0–9.8 mm);
(2) blackish pigments on the entire dorsum of head (Figure 27);
278
C. Molineri and D. Emmerich
Figures 30–34. Campsurus amapaensis sp. nov.; (30) ventral view; (31) lateral view; (32) tip of
penis lobe, lateral view; (33) tip of left penis lobe, ventral view; (34) detail of pedestal, ventral
view.
(3) pronotum shaded greyish, blackish medial line present Figure 27);
(4) abdominal colour pattern strongly marked (Figures 28 and 29), pale
transverse dashes on terga II–VII broken at middle and forming pale
submedian circles (mainly on V–VII);
(5) pedestals narrow, outer posterolateral margin reduced, inner margin rounded
but with acute apex (Figure 34);
(6) penes moderately twisted, similar to C. argentinus but with a more acute apex
(Figures 32 and 33).
Campsurus amapaenis sp. nov. is very similar to C. argentinus, sharing some
features of colour pattern (head and thorax) and form of penes, but the new species
can easily be distinguished because of their narrow pedestals (Figure 34) and
abdominal colour pattern (Figures 28 and 29).
Distribution
Known only from the type locality, in northeast Brazil (Amapá).
Aquatic Insects
279
Figures 35–40. Male wings; (35–36) C. argentinus, (35) forewing; (36) hind wing; (37–38) C.
major, (37) forewing; (38) hind wing; (39–40) C. amapaensis sp. nov.; (39) forewing; (40) hind
wing.
Acknowledgements
We are grateful to Janice G. Peters (FAMU) for permanent help and support during the
course of this study. Thanks also to Lucimar Gomes Dias and Iracenir Santos Andrade for the
loan of material from Amapá. Richard Hoebeke and James Liebher kindly attended our
interest in studying the types at CUIC. Financial support from Pedeciba-Biologı́a (Naciones
Unidas Proyecto URU/06/004), CONICET (PIP 1484) and ANPCYT (PICT 528, 351) is
greatly acknowledged.
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